La spécialisation des parasites vis-à-vis de leur(s) hôte(s) et la formation de races d'hôtes sont des processus évolutifs clés dans le maintien et l'émergence de la diversité au sein des populations de parasites. Notre objectif était d'étudier ces processus chez Ixodes uriae, une tique d'oiseaux marins présentant une vaste distribution géographique et une large diversité d'hôtes, afin de mieux comprendre le rôle relatif des contraintes liées à l'hôte et aux facteurs géographiques dans l'évolution et la diversification de ce parasite. Cette question a été abordée à travers différentes approches réunissant des analyses de génétique des populations et de phylogéographie, des analyses morphologiques et une approche expérimentale sur le terrain. Les résultats obtenus montrent que le facteur spatial joue un rôle important dans la diversification d'I. uriae puisque quatre grands groupes géographiques génétiquement isolés ont été identifiés. L'évolution de races d'hôtes est également un processus récurrent de l'évolution d'I. uriae, même si la divergence entre races est plus ou moins marquée d'une région à l'autre. L'évolution des races semble impliquer une préférence d'hôte chez la tique ainsi que des contraintes adaptatives liées à l'hôte, de type mécanique ou physiologique (e.g., bec des oiseaux, digestion du sang, réponse immunitaire). L'ensemble des résultats est cohérent avec la notion d'évolution en mosaïque géographique qui prédit que les interactions entre espèces peuvent évoluer de manière différente dans un contexte spatial hétérogène ; ce qui souligne l'importance des caractéristiques écologiques des hôtes dans la diversification de la tique I. uriae. D'après nos résultats et ceux de différents collègues, la spécialisation d'hôte pourrait s'avérer être un processus commun chez les tiques et donc avoir de profondes implications épidémiologiques et évolutives pour les pathogènes qu'elles transmettent. / Host specialization and host-race formation may be key processes in maintaining and generating diversity within parasite populations. The objective of this thesis was to analyze these processes in the seabird tick Ixodes uriae, a common and widely distributed ectoparasite that exploits a great diversity of seabird host species, to better understand the relative role of host-related constraints and geographic factors in its evolution and diversification. I addressed this question using different approaches, including population genetic and phylogeographic analyses, morphometry and field experimentation. Results show that the spatial factor is a key component in shaping the diversity of I. uriae, as four geographical genetically isolated groups exist within the global distribution of this parasite. The formation of host races is also a general process in the evolution of I. uriae, but has evolved to different degrees in different geographical regions. Host preference in the tick, along with mechanical or physiological host-related constraints (e.g., beak morphology, blood digestion, immune response) are likely involved in the evolution of I. uriae host races. Overall, results fall in line with the geographic mosaic theory of evolution that predicts that the outcome of species interactions can vary across geographic landscapes, highlighting the role played by ecological characteristics of the host in the diversification of I. uriae. Given results to date, host specialization may be a common process in tick systems and thus may have strong epidemiological and evolutionary implications for the pathogens that they transmit.

Orientador: Edson Aparecido Adriano / Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-26T12:26:07Z (GMT). No. of bitstreams: 1 Naldoni_Juliana_D.pdf: 10311500 bytes, checksum: a497c164e89d09cfdbe8ccfcf87ef274 (MD5) Previous issue date: 2014 / Resumo: Mixosporídeos são cosmopolitas e infectam peixes em diversas regiões do mundo. Atualmente são conhecidas cerca de 2.400 espécies, das quais a grande maioria é parasito de peixes, tanto de ambiente natural como de sistemas de criação, sendo algumas espécies responsáveis por altas taxas de mortalidade em várias partes do mundo. Este trabalho teve como objetivo o estudo da diversidade de mixosporídeos parasitos de Pseudoplatystoma corruscans (pintado), Salminus franciscanus (dourado) e Brycon orthotaenia (matrinxã) da bacia do rio são Francisco, município de Pirapora, MG, Brasil. Foram realizados estudos morfológicos, ultraestruturais, histotológicos e moleculares de cinco novas espécies de mixosporídeos, sendo uma do gênero Henneguya em pintado, duas do gênero Myxobolus infectando dourado e duas infectando matrinxã. Henneguya sp. n. 1 apresentou plasmódios brancos e alongados no tecido conjuntivo das brânquias de pintado. A análise ultraestrutual revelou a parede plasmodial com delicadas projeções em direção aos tecidos do hospedeiro e a presença de uma fina camada de material finamente granular isolando o parasito do contato com o tecido do hospedeiro. A análise histológica revelou que o desenvolvimento do plasmódio causou a compressão no tecido conjuntivo e epitelial, deformação dos filamentos e a fusão lamelar. A análise filogenética, baseada no gene 18S rDNA e utilizando somente espécies dos gêneros Henneguya e Myxobolus parasitos de siluriformes, revelou o agrupamento de acordo com a família dos peixes hospedeiros. Myxobolus sp. n. 1 apresentou plasmódios brancos e arredondados entre os raios da nadadeira de dourado. A análise ultraestrutural revelou uma camada de fibroblastos circundando o plasmódio, impedindo o contato com as células do hospedeiro. Myxobolus sp. n. 2 apresentou plasmódios brancos e arredondados no fígado de dourado. Myxobolus sp. n. 3 e Myxobolus sp. n. 4 apresentaram plasmódios brancos e arredondados, sendo que a primeira ocorreu no baço e a segunda no rim de matrinxã. A análise ultraestrutural de Myxobolus sp. n. 2, Myxobolus sp. n. 3 e Myxobolus sp. n. 4 revelou o contato direto entre a parede do plasmódio do parasito e o tecido dos hospedeiros. As paredes dos plasmódios das cinco espécies aqui estudadas foram compostas por membrana simples. O processo de esporogênese das cinco espécies foi assincrônico, com células germinativas e jovens estágios de desenvolvimento dos esporos ocorrendo na periferia do plasmódio e esporos imaturos e maduros foram observados na região central. A análise filogenética, baseado no gene 18S rDNA e usando somente espécies dos gêneros Henneguya e Myxobolus parasitos de peixes da América do Sul mais as quatro novas espécies de Myxobolus parasitas de briconídeos, mostrou a especificidade de hospedeiro e a afinidade de órgão/tecido, como um importante sinal evolutivo para Myxobolus/Henneguya / Abstract: Myxosporeans are cosmopolitan parasites and infect fish in various regions of the world. So far, are known about 2,400 species, of which the vast majority are parasites of fishes, from natural environment and fish farms, and some species responsible for high mortality rates in various parts of the world. This work aimed to study the diversity of myxosporeans of Pseudoplatystoma corruscans (pintado), Salminus franciscanus (dourado) and Brycon orthotaenia (matrinxã) from the São Francisco River, municipality of Pirapora, MG, Brazil. Morphological, ultrastructural, histological and molecular studies of five new species of myxosporeans were performed, being one species of the genus Henneguya infecting pintado, two of the genus Myxobolus infecting dourado and two infecting matrinxã. Henneguya sp. n. 1 had white and elongated plasmodia in the connective tissue of the gill filaments. The ultrastructural analysis revealed the plasmodial wall with delicate projections towards the tissues of the host, and the presence of a thin layer of fibrous material isolating the parasite of the contact with the host tissue. Histological analysis revealed that the development of the plasmodium caused compression of the connective and epithelial tissue, deformation of the filament and lamellar fusion. Phylogenetic analysis, based on 18S rDNA gene, and using only Henneguya and Myxobolus parasites of siluriformes revealed clustering according to the family of the host fish. Myxobolus sp. n. 1 had white and rounded plasmodia that developed between the fin rays of dourado. The ultrastructural analysis showed a fibroblast layer surrounding the plasmodium, preventing contact of the parasite with the host tissues. Myxobolus sp. n. 2 had white and rounded plasmodia that developed in the liver also of dourado. Myxobolus sp. n. 3 and Myxobolus sp. n. 4 infected matrinxã, being that the first had white and rounded plasmodia in the spleen and the second in the kidney. The ultrastructural analyses of Myxobolus sp. n. 2, Myxobolus sp. n. 3 and Myxobolus sp. n. 4 revealed direct contact between the plasmodial wall and the host tissue. The plasmodial wall of the five myxosporeans species subject of this study was composed by single membrane. The process of sporogenesis in these five species was asynchronous, with germ cells and young development stages of spores occurring in the periphery of the plasmodia and immature and mature spores in the central region. Phylogenetic analysis based on 18S rDNA gene and using only Henneguya and Myxobolus parasites of fish from South America plus the four new Myxobolus species parasites of bryconids, shows host specificity and organs/tissue affinity as important evolutionary signs to Myxobolus/Henneguya / Doutorado / Parasitologia / Doutora em Parasitologia

Schistosoma mansoni é um dos agentes causadores da esquistossomose, doença infecciosa negligenciada que afeta milhões de pessoas no mundo. É um platelminto parasitário dióico, com um complexo ciclo de vida, composto de seis estágios. Nos últimos cinco anos, projetos de seqüenciamento em larga escala de etiquetas de genes expressos (ESTs) de Schistosoma geraram uma quantidade razoável de dados que ainda pode ser mais bem explorada. O objetivo central deste trabalho é analisar computacionalmente a expressão gênica de S. mansoni, sob três focos distintos: (i) micro-arranjos de DNA, para os quais descrevemos o desenho e análise de dados de uma plataforma de cDNA (4,600 elementos) e outra de oligonucleotídeos (44,000 elementos). Estão também descritas diversas ferramentas de análise que implementamos e são amplamente usadas em nosso grupo. Os micro-arranjos têm servido de base para vários projetos, como o estudo da resposta do parasita a hormônios e drogas e expressão gênica durante o ciclo de vida. (ii) Identificação in silico de pares de transcritos senso- antisenso com possível ação em trans, potencialmente importantes na regulação gênica. (iii) Análises da coleção de ESTs existentes sob uma perspectiva evolutiva. Através dessa abordagem encontramos genes importantes como um possível inibidor de angiogênese e um regulador da via do mevalonato, conhecido como essencial para a produção de ovos; estes constituem a principal causa de morbidez da esquistossomose. Os resultados aqui apresentados contribuem para o entendimento da complexa biologia inerente ao ciclo de vida de S. mansoni e para acelerar a busca de futuras possibilidades de tratamento. / Schistosoma mansoni is one of the causative agents of schistosomiasis, a neglected infectious disease which affects millions of people worldwide. It is a dioecious parasitic platyhelminth, with a complex life cycle composed of six stages. In the past five years, large scale sequencing projects have generated a reasonable amount of expressed sequence tag (EST) data that can still be better explored. The goal of this thesis is to computationally analyze the S. mansoni gene expression, under three different focuses: (i) DNA microarrays, for which we describe the design and data analyses of a cDNA (4,600 elements) and an oligonucleotide (44,000 elements) platform. We also describe the implementation of several analysis tools which are widely used in our group. Our microarrays are being used in several projects, such as the study of parasite response to drugs and hormones, as well as its gene expression pattern during the life cycle. (ii) In silico identification of possible trans acting natural sense-antisense pairs, potentially important in gene regulation. (iii) Analyses of the available EST dataset under an evolutionary perspective. We have found interesting genes such as a possible angiogenesis inhibitor and a regulator of the mevalonate pathway, known to be essential for egg production; eggs are the main cause of morbidity of schistosomiasis. The results reported here contribute to the understanding of the complex biology underlying the S. mansoni life cycle and to accelerate the search for future possibilities of treatment.

Une caractéristique majeure de l’infection par Toxoplasma gondii est le contrôle rapide de la population parasitaire par une réponse immunitaire engageant des cellules résidentes et recrutées ainsi que des cytokines pro- et anti-inflammatoire. Dans ce contexte, l’IFNγ active une multitude d’activité anti- T. gondii des cellules immunes et non-immunes, mais peut aussi contribuer à l’immunopathologie. T. gondii a élaboré des mécanismes pour contrer les défenses de l’hôte en interférant avec la transcription des gènes stimulés par l’IFNγ. Nous avons identifié TgIST (T. gondii inhibitor of STAT1 transcriptional activity) comme un interrupteur moléculaire exporté par les parasites intracellulaires et qui est localisé dans le noyau des cellules hôtes, où il inhibe l’expression des gènes pro-inflammatoires dépendants de STAT1. Nous avons montré que TgIST séquestre STAT1 à des sites spécifiques, et promeut la formation de chromatine non permissive grâce à sa capacité à recruter le remodeleur chromatinien NuRD. Nous avons montré que durant l’infection aiguë en souris, les parasites déficients pour TgIST sont rapidement éliminés par les monocytes pro-inflammatoires GR1+, ce qui montre le rôle protecteur de TgIST contre les défenses médiées par l’IFNγ. En révélant les fonctions de TgIST, cette étude montre de nouvelles évidences sur la façon dont T.gondii a élaboré une arme moléculaire de choix pour prendre le contrôle sur la réponse immune, de façon à promouvoir le parasitisme à long terme / An early hallmark of Toxoplasma gondii infection is the rapid control of the parasite population by a potent multifaceted innate immune response that engages resident and homing immune cells along with pro- and counter-inflammatory cytokines. In this context, IFN-γ activates a variety of T. gondii–targeting activities in immune and nonimmune cells but can also con- tribute to host immune pathology. T. gondii has evolved mechanisms to timely counteract the host IFN-γ defenses by interfering with the transcription of IFN-γ–stimulated genes. We now have identified TgIST (T. gondii inhibitor of STAT1 transcriptional activity) as a critical molecular switch that is secreted by intracellular parasites and traffics to the host cell nucleus where it inhibits STAT1-dependent proinflammatory gene expression. We show that TgIST not only sequesters STAT1 on dedicated loci but also promotes shaping of a nonpermissive chromatin through its capacity to recruit the nucleosome remodeling deacetylase (NuRD) transcriptional repressor. We found that during mice acute infection, TgIST-deficient parasites are rapidly eliminated by the homing Gr1+ inflammatory monocytes, thus highlighting the protective role of TgIST against IFN-γ–mediated killing. By uncovering TgIST functions, this study brings novel evidence on how T. gondii has devised a molecular weapon of choice to take control over a ubiquitous immune gene expression mechanism in metazoans, as a way to promote long-term parasitism.

<p> Previous studies dealing with the structure and function of the attachment organs in the strigeid trematodes have neglected to describe the processes involved in the formation of attachment. A knowledge of these processes is necessary to promote the understanding of the host-parasite relationship. </p> <p> In the present study, specimens of developing Cotylurus variegatus were examined using light and electron microscopic techniques. It seemed relevant to consider not only the sequence of attachment events, but the growth and structure of the attachment organs in relation to the total parasite body growth and structure. This of course, has led to considerations of the reputative functions of these structures. </p> / Thesis / Master of Science (MSc)

Cuscuta pentagona is an obligate parasitic plant that hinders production of crops throughout the world. Parasitic plants have unique morphological and physiological features, the most prominent being the haustorium, a specialized organ that functions to connect them with their host's vascular system. The Cuscuta haustorium is remarkable in that it enables mRNA movement to occur between hosts and parasite, but little is known about the mechanisms regulating cross-species mRNA transfer or its biological significance to the parasite. These questions were addressed with genomics approaches that used high throughput sequencing to assess the presence of host mRNAs in the parasite as well as parasite mRNAs in the host. For the main experiment Cuscuta was grown on stems of Arabidopsis thaliana and tomato (Solanum lycopersicon) hosts because the completely sequenced genomes of these plants facilitates identification of host and parasite transcripts in mixed mRNA samples. Tissues sequenced included the Cuscuta stem alone, the region of Cuscuta-host attachment, and the host stem adjacent to the attachment site. The sequences generated from each tissue were mapped to host reference genes to distinguish host sequences, and the remaining sequences were used in a de novo assembly of a Cuscuta transcriptome. This analysis revealed that thousands of different Arabidopsis transcripts, representing nearly half of the expressed transcriptome of Arabidopsis, were represented in the attached Cuscuta. RNA movement was also found to be bidirectional, with a substantial proportion of expressed Cuscuta transcripts found in host tissue. The mechanism underlying the exchange remains unknown, as well as the function of mobile RNAs in either the parasite or host. An approach was developed to assay potential translation of host mRNAs by detecting them in the Cuscuta translatome as revealed by sequencing polysomal RNA and ribosome-protected RNA. This work highlights RNA trafficking as a potentially important new form of interaction between hosts and Cuscuta. / Ph. D.

Co-evolution between parasites and their hosts result in extremely well-orchestrated and intimate relationships that are characterized by remarkable adaptations in the attack response of the parasite and the defense response of the host. To fully understand host-parasite interactions, these adaptations must be considered in the context of the ecological constraints in which they evolved. As a serious pest to salmon mariculture, Lepeophtheirus salmonis has been extensively studied; however, there are still several areas that require further research. Of utmost importance, and the topic of this thesis, is molecular basis for resistance to sea lice. The following chapters investigate this phenomena under the umbrella of ecological immunology using combined modern technologies of transcriptomics, proteomics and functional immunology with a focus on the primary interaction site. In the first chapter, I describe the key players involved in this host-parasite relationship with a focus on the primary interaction site, the louse-salmon interface, where there are responses by the louse (attack) and the salmon host (defense). Previous research indicated that an early aggressive inflammatory response at the louse-skin interface contributes to resistance in coho salmon; however, there are no data characterizing a site-specific response in resistant (pink and coho) and susceptible (Atlantic, chum) species. Accordingly in Chapter 2, I define site-specific cutaneous responses in Atlantic, pink and chum salmon to establish genetic biomarkers of resistance. Chapter 3 focuses on identification of cellular effectors using histochemical localization of biomarkers to characterize cellular populations activated at the louse-attachment site, while broadening the gene targets. Our notion of pink salmon as a resistant species is challenged by the common observation of migrating pink salmon supporting large populations of L. salmonis in the field. Thus the purpose of chapter 4 was to investigate potential mechanisms to explain variations in susceptibility as a function of life history. Host-parasite relationships are a product of both host and parasite responses; therefore, in chapters 5 and 6, I shift focus to the level of the parasite. In chapter 5 I present the first documented large-scale transcriptomic profiling of L. salmonis during feeding on both resistant (coho) and susceptible (Atlantic, sockeye) salmon. This was followed (chapter 6) by describing the proteomic profile of L. salmonis secretions after feeding on Atlantic salmon. In the seventh and final chapter, I present my conclusions on the molecular mechanisms for resistance to sea lice and discuss potential applications of this information for future louse control strategies. / Graduate

Les parasites exercent une pression de sélection quasiment universelle. Cette thèse aborde les relations hôte-parasite dans le système Lymnaeidae (Gastropoda) Fasciolidae (Trematoda, douves) sous différents aspects, afin de brosser une image de cette interaction et de son évolution. J'ai tout d'abord établi les relations phylogénétiques entre les espèces de Lymnaeidae, puis retracé l'évolution de différents caractères, tels que la susceptibilité à l'infestation par Fasciola hepatica et F. gigantica. Alors que F. hepatica est un parasite généraliste, capable d'infester des mollusques de presque tous les clades de la famille Lymnaeidae, l'infestation par F. gigantica est plus restreinte à un clade. J'ai ensuite étudié plus finement la coévolution entre le parasite F. hepatica et deux de ses hôtes Lymnaeidae (Galba truncatula et Omphiscola glabra) au sein d'une métapopulation, ce qui a confirmé la stratégie généraliste de F. hepatica. En plus, il semblerait que les mollusques parasités et non parasités de G. truncatula aient des différences génétiques, au moins dans cinq des huit populations étudiées. J'ai caractérisé la diversité génétique de deux espèces de mollusques envahissantes, préférentiellement autogames et impliquées dans la transmission de F. hepatica : Pseudosuccinea columella et Lymnaea sp. On trouve une diversité génétique très réduite, chez ces deux espèces, ce qui pourrait faciliter leur expansion géographique et leur infestation par F. hepatica. Ce travail m'a ensuite conduit à mesurer le temps d'attente avant l'autofécondation et la dépression de consanguinité chez ces deux espèces. J'ai trouvé que ces deux espèces sont caractérisées par une dépression de consanguinité très faible et un temps d'attente nul, ce qui confirme les résultats obtenus lors d'une collaboration dans une étude à plus large échelle. Cette thèse souligne l'importance des études en évolution pour comprendre l'épidémiologie des maladies parasitaires. / Parasites constitute a selective pressure to almost all living beings. This thesis addresses the host-parasite interaction in the Lymnaeidae (Gastropoda) Fasciolidae (Trematoda; liver flukes) system through different approaches, with the aim to give a comprehensive image of this interaction and its evolution. First, I established the phylogenetic relationships among Lymnaeidae species, and then mapped the evolution of different characters such as the susceptibility to the infection by Fasciola hepatica and F. gigantica. While F. hepatica is a generalist parasite, capable to infect snails from almost all clades of the Lymnaeidae, infection by F. gigantica is restricted to one clade. Next, I studied the co-evolution between the parasite F. hepatica and two of its intermediate host species (Galba truncatula and Omphiscola glabra) at a finer scale: within a metapopulation. This study confirmed the generalist strategy of F. hepatica. In addition, it seems that parasitized and non-parasitized G. truncatula snails exhibit genetic differences, at least in five out of eight studied populations.I also characterized the genetic diversity of two species of invasive snails involved in the transmission of F. hepatica: Pseudosuccinea columella and Lymnaea sp. We discuss the possible reasons of invasion success in these snails, despite their low genetic diversity, which could facilitate their infection by F. hepatica. Their capacity to respond to parasitism is certainly reduced, all the more that these species are preferential selfers. This work has then led me to measure the waiting time before self-fertilization and inbreeding depression in these two snails. I found that these two species are characterized by low inbreeding depression and present no waiting time, which confirms the results obtained in a collaborative project at larger phylogenetic scale. This thesis strengthens the importance of evolutionary studies to understand the epidemiology of parasitic diseases.

This thesis focuses on mechanisms and processes underlying spatial patterns of resistance and virulence and on local adaptations in plant–parasite interactions. The model system used comprises the plant host Salix triandra, the pathogenic rust fungus Melampsora amygdalinae, the leaf beetle Gonioctena linnaeana, and the galler Pontania triandrae. In this work, I (1) emphasize the most important factors determining the outcome of a plant–pathogen interaction, and the types of systems in which local adaptations can be expected, (2) examine the resistance structures of different populations of S. triandra, and whether the leaf beetle G. linnaeana responds to the local conditions of the populations of S. triandra in Sweden, and (3) address whether the distribution of parasites on S. triandra can be explained by the plant content of secondary metabolites. A review of several studies of the subject leads to the conclusion that adaptation of pathogens to their local hosts is more likely to be found in systems in which the pathogen is host-specific, non-systemic, and has a larger dispersal range and evolutionary potential than its host does. Furthermore, the scale of the study must be adjusted to that of the pathogen’s local population distribution. In addition, the temporary nature of host–pathogen interactions influences the importance of sample size, and too-small sample sizes can lower the chance of finding local adaptations, even though they may have evolved in a given system. The results of an inoculation experiment using material from physically isolated natural populations of S. triandra and M. amygdalinae confirm the importance of previous conclusions. Spatial variation in the resistance structure of S. triandra also has effects on the insect herbivore G. linnaeana, which has responded by adapting to the local hosts. However, local differences in secondary chemistry affect different parasites in different ways, and while P. triandrae is attracted by high levels of phenolic compounds, including tannins, M. amygdalinae and G. linnaeana are more rarely found on plant individuals with high concentrations of tannins. In addition, brood deposition by adult females of G. linnaeana and the performance of larvae are positively affected by luteolin-7-glucoside and an additional unidentified flavonoid, whereas they are negatively affected by the presence of (+)-catechin and high levels of tannins. Our results also show that plants traits that provide resistance to one type of parasite do not necessarily provide resistance to others. This indicates that different natural enemies potentially assert divergent selection pressure on S. triandra phenotypes which can be important for maintaining phenotypic variation in plant species.

L’expression du phénotype des organismes dépend en partie d’organismes symbiotiques avec qui ils sont en interaction étroite. Selon le mode de transmission du symbiote, ce dernier va être en conflit d’intérêt plus ou moins intense avec l’hôte pour l’expression du phénotype, conduisant parfois le symbiote à évoluer vers la manipulation du phénotype de l’hôte. Nous avons tenté d’identifier différents facteurs génétiques et environnementaux influençant l’expression et l’évolution de la manipulation chez l’insecte parasitoïde de larves de drosophiles, Leptopilina boulardi, et son virus manipulateur du comportement, LbFV. Ce virus bénéficie d’une transmission mixte, verticale et horizontale, cette dernière étant favorisée par l’induction de superparasitisme induite par le virus. L’étude de la contribution du génotype du parasitoïde dans l’expression de la manipulation a révélé la présence de gènes de résistance partielle à la manipulation. Le potentiel évolutif de cette résistance a ensuite été évalué par des expériences d’évolution expérimentale. Nous avons également montré que LbFV augmente la virulence du parasitoïde envers les larves de drosophiles, révélant ainsi une évolution vers une forme de mutualisme sur ce trait. Par ailleurs, le travail montre qu’un même parasitoïde peut être non seulement infecté par plusieurs souches du virus LbFV mais également infecté par un virus à ARN, décrit pour la première fois dans cette thèse. La transmission verticale, la prévalence élevée et les forts effets phénotypiques de ce virus soulignent de nouveau l’importance des virus dans l’expression du phénotype en population naturelle. / The expression of the phenotype of an individual depends partly on the presence of symbiotic organisms with which it engages in intimate interactions. According to the symbiont’s mode of transmission, the conflict of interest between the host and its symbiont for the expression of the phenotype can be more or less stronger, sometimes leading to the evolution of manipulation of the host phenotype by the symbiont. We attempted to identify different genetic and environmental factors influencing the expression and the evolution of manipulation in the Drosophila parasitoid wasp, Leptopilina boulardi, and its behavior manipulating virus, LbFV. The virus undergoes both vertical and horizontal transmission, the latter being favoured by the induction of superparasitism behaviour by the virus. The study of the contribution of the parasitoid genotype to the expression of the manipulation revealed a polymorphism in the resistance to the manipulation. The evolutionary potential of this resistance was then investigated using a protocol of experimental evolution. We also demonstrated that LbFV increases the virulence of the parasitoid towards its Drosophila host, revealing a form of mutualistic interaction on this trait. Additionally, we showed that an individual parasitoid can be coinfected by several LbFV strains but also by an RNA virus, described for the first time in this thesis. The vertical transmission, the high prevalence and the strong phenotypic effects of this new virus further highlights the importance of viruses in the expression of the phenotype in natural populations.